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Univerzita Palackého V Olomouci Přírodovědecká UNIVERZITA PALACKÉHO V OLOMOUCI PŘÍRODOVĚDECKÁ FAKULTA KATEDRA ZOOLOGIE MOLEKULARNÍ IDENTIFIKACE DRUHOVÉ DIVERSITY RODU AGRILUS (COLEOPTERA: BUPRESTIDAE) Diplomová práce Vypracovala: Bc. Ivana Kelnarová Studijní program: Biologie Studijní obor: Zoologie Forma studia: prezenční Školitel: Prof. Ing. Ladislav Bocák, Ph. D. Olomouc 2017 PROHLÁŠENÍ Prohlašuji, že jsem bakalářkou práci vypracovala samostatně pod vedením vedoucího Prof. Ing. Ladislava Bocáka Ph. D. a použila jsem pouze uvedené bibliografické zdroje. V Olomouci dne 28. 7. 2017 ………………………… PODĚKOVÁNÍ Ráda bych tímto poděkovala Prof. Ing. Ladislavu Bocákovi, Ph.D za odborné konzultace k této práci. Taktéž musí poděkovat Mgr. Renatě Bílkové za pomoc při práci v laboratoři. Za podporu velmi děkuji svému příteli, rodině a kamarádům. BIBLIOGRAFICKÁ IDENTIFIKACE Jméno a příjmení autora: Bc. Ivana Kelnarová Název práce: Molekularní identifikace druhové diversity rodu Agrilus (Coleoptera: Buprestidae) Typ práce: Diplomová práce Pracoviště: Katedra zoologie, Přírodovědecká fakulta, Olomouc Vedoucí práce: Prof. Ing. Ladislav Bocák Ph. D. Rok obhajoby práce: 2017 ABSTRAKT: Tato studie se zabývá klasifikací rodu Agrilus (Coleoptera: Buprestidae), nejpočetnější, značně morfologicky uniformní skupiny hmyzu, která obsahuje více než 3000 druhů. Mnohé z těchto druhů jsou významnými škůdci, kteří se za příhodných podmínek stávají v některých oblastech invazivními. Značná morfologická podobnost jednotlivých druhů znesnadňuje jejich jednoznačnou identifikaci, která je pro rychlé zachycení výskytu neznámého škůdce nezbytná. Určit neznámý druh rodu Agrilus, zvláště pokud je nepůvodní, je bez rozsáhlé srovnávací sbírky téměř nemožné a specialistů zabývajících se rodem Agrilus je velmi málo. Tato práce nabízí možnost identifikace neznámých vzorků pomocí databáze molekulárních markerů zahrnující téměř 100 druhů rodu Agrilus, jež je dostupná v databázi GenBank a použitelná pro molekulárně založenou identifikaci nových vzorků. V této studii byly použity dva přístupy: byly vytvořeny fylogenetické stromy na základě molekulárních markerů s použitím metody maximum likelihood a dále byly srovnány genetické párové vzdálenosti. Analýza 'barcode' fragmentu neumožňuje identifikace, pokud daný druh již není zachycen v databázi. V případě analýzy více fragmentů, je možné po nenalezení odpovídajícího jedince pomocí fylogenetické hypotézy o příbuznosti druhů nalézt nejpříbuznější sesterskou linii s jistým geografickým původem a odhadnout tak původ neznámého vzorku. V databázi jsou rovněž zachyceny nepůvodní druhy zavlečené do Severní Ameriky z Evropy a Asie. Klíčová slova: Agrilus, invazivní druhy, databáze DNA markerů, barcode, cox1 mtDNA, rrnL mtDNA, RAxML Počet stran: 51 Počet stran příloh: 31 Jazyk: český BIBLIOGRAPHIC IDENTIFICATION First name and surname of the author: Bc. Ivana Kelnarová Name of the thesis: Molecular identification of specific diversity in Agrilus (Coleoptera: Buprestidae) Type of thesis: Diploma thesis Workplace: Department of Zoology, Faculty of Science, Olomouc Supervisor: Prof. Ing. Ladislav Bocák Ph. D. Year of defense: 2017 ABSTRACT: This work tests the methods for DNA-based identification of the largest, morphologically uniform insect lineage. The genus Agrilus (Coleoptera: Buprestidae), which includes more than 3000 morphologically identified species, contains economically important pests, which become invasive in some areas with suitable conditions. The unambiguous identification, which is necessery for quick interception and disposal, is complicated by high morphological similarity of individual species. The identification of the unknown species of Agrilus is impossible without an extensive reference collection and expertize knowledge and additionally, there are few trained entomologists available. This study demonstrates the effectiveness of various methods for identification of unknown samples. I used a newly produced molecular marker database, which represents about 100 species of the genus Agrilus from the Northern Hemisphere and the data available in GenBank database. The tree-based and distance based methods were used for identification of species limits and estimation of a cryptic genetic diversity. In this study, phylogenetic hypotheses were also constructed based using molecular markers and maximum likelihood optimality criterion. If the closely similar sequence is absent in the database, to data can identify the most closely related sister species of population, inable estimation of the geographic origin based on the phylogenetic hypothesis. Key words: Agrilus, Invasive species, DNA markers database, barcode, cox1 mtDNA, rrnL mtDNA, RAxML Number of pages: 51 Number of appendices: 31 Language: Czech Obsah 1 ÚVOD ............................................................................................................................... 7 2 INVAZIVNÍ DRUHY ...................................................................................................... 9 2.1 Vymezení pojmu 'invazivní druh' ............................................................................ 9 2.2 Zavlečení nepůvodního druhu, jeho invazivita a vliv na původní flóru a faunu ..... 9 2.3 Invazivní druhy řádu Coleoptera............................................................................ 10 2.3.1 Rod Agrilus a invazivita jeho druhů .................................................................. 12 2.4 Opatření proti šíření invazivních druhů ................................................................. 16 3 CÍLE PRÁCE .................................................................................................................. 18 4 METODIKA ................................................................................................................... 19 4.1 Materiál .................................................................................................................. 19 4.2 Laboratorní protokoly ............................................................................................ 19 4.2.1 Izolace DNA ...................................................................................................... 19 4.2.2 PCR amplifikace DNA ....................................................................................... 20 4.2.3 Elektroforéza a čištění PCR produktu ................................................................ 21 4.2.4 Sekvenační reakce .............................................................................................. 22 4.2.5 Čištění produktu sekvenační reakce ................................................................... 22 4.2.6 Sekvenování ....................................................................................................... 23 4.3 Analýza DNA sekvencí a fylogenetická analýza ................................................... 23 4.4 Delimitace druhů .................................................................................................... 23 4.4.1 Species identifier ................................................................................................ 24 4.4.2 bPTP ................................................................................................................... 24 4.5 Analýza statické podpory pro zjištěné fylogenetické vztahy ................................. 24 4.6 Vyhodnocení geografického původu fylogeneticky příbuzných druhů ................. 24 5 VÝSLEDKY ................................................................................................................... 25 5.1 Fylogenetické vztahy ............................................................................................. 26 5.2 Molekulární delimitace druhů ................................................................................ 29 5.3 Použité zkratky ....................................................................................................... 30 6 DISKUZE A ZÁVĚR ..................................................................................................... 37 6.1 Srovnání morfologicky a molekulárně určených druhů ......................................... 38 7 SEZNAM POUŽITÉ LITERATURY ............................................................................ 42 8 SEZNAM PŘÍLOH ......................................................................................................... 51 1 ÚVOD Krasci rodu Agrilus Curtis, 1825 (Coleoptera: Elateriformia: Buprestidae) představují svým počtem cca 3000 druhů nejpočetnější rod živočichů v celé živočišné říši. Jejich rozšíření je kosmopolitní a největší diversita je v oblastech s vysokou rozmanitostí potenciálních hostitelských rostlin, především v subtropických částech holoarktické oblasti (Bellamy 2010). Všechny druhy této skupiny jsou fytofágní a jejich diversita je pravděpodobně výsledkem četných přesunů z jedné hostitelské rostliny na druhou. Příkladem druhu, jehož diversifikace je ovlivněna přesuny mezi hostitelskými rostlinami je polyfágní středoevropský druh Agrilus viridis (Bernhard et al. 2005). Analýza molekulárních dat tohoto druhu získaného z odlišných živných rostlin naznačuje, že by se ve skutečnosti mohlo jednat o více specializovaných populací, poddruhů nebo druhů, a nikoliv pouze o jediný takto široce polyfágní druh (Bernhard et al. 2005, Pentinsaari et al. 2014). Larvy rodu Agrilus se živí rostlinnými pletivy a daří se jim především pod kůrou rozmanitých druhů stromů a keřů (Jendek 1994). Větší počty jedinců se vyvíjejí na jednotlivých rostlinách zejména v případě, že jsou stresovány suchem (Dunn
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